Building behavior does not drive rates of phenotypic evolution in spiders.

Do animals set the course for the evolution of their lineage when manipulating their environment? This heavily disputed question is empirically unexplored but critical to interpret phenotypic diversity. Here, we tested whether the macroevolutionary rates of body morphology correlate with the use of built artifacts in a megadiverse clade comprising builders and nonbuilders-spiders. By separating the inferred building-dependent rates from background effects, we found that variation in the evolution of morphology is poorly explained by artifact use. Thus natural selection acting directly on body morphology rather than indirectly via construction behavior is the dominant driver of phenotypic diversity.

Diet assessment of two land planarian species using high-throughput sequencing data.

Geoplanidae (Platyhelminthes: Tricladida) feed on soil invertebrates. Observations of their predatory behavior in nature are scarce, and most of the information has been obtained from food preference experiments. Although these experiments are based on a wide variety of prey, this catalog is often far from being representative of the fauna present in the natural habitat of planarians. As some geoplanid species have recently become invasive, obtaining accurate knowledge about their feeding habits is crucial for the development of plans to control and prevent their expansion. Using high throughput sequencing data, we perform a metagenomic analysis to identify the in situ diet of two endemic and codistributed species of geoplanids from the Brazilian Atlantic Forest: Imbira marcusi and Cephaloflexa bergi. We have tested four different methods of taxonomic assignment and find that phylogenetic-based assignment methods outperform those based on similarity. The results show that the diet of I. marcusi is restricted to earthworms, whereas C. bergi preys on spiders, harvestmen, woodlice, grasshoppers, Hymenoptera, Lepidoptera and possibly other geoplanids. Furthermore, both species change their feeding habits among the different sample locations. In conclusion, the integration of metagenomics with phylogenetics should be considered when establishing studies on the feeding habits of invertebrates.

The effect of fight cost structure on fighting behaviour involving simultaneous decisions and variable investment levels.

In the "producer-scrounger" model, a producer discovers a resource and is in turn discovered by a second individual, the scrounger, who attempts to steal it. This resource can be food or a territory, and in some situations, potentially divisible. In a previous paper we considered a producer and scrounger competing for an indivisible resource, where each individual could choose the level of energy that they would invest in the contest. The higher the investment, the higher the probability of success, but also the higher the costs incurred in the contest. In that paper decisions were sequential with the scrounger choosing their strategy before the producer. In this paper we consider a version of the game where decisions are made simultaneously. For the same cost functions as before, we analyse this case in detail, and then make comparisons between the two cases. Finally we discuss some real examples with potentially variable and asymmetric energetic investments, including intraspecific contests amongst spiders and amongst parasitoid wasps. In the case of the spiders, detailed estimates of energetic expenditure are available which demonstrate the asymmetric values assumed in our models. For the wasps the value of the resource can affect the probabilities of success of the defender and attacker, and differential energetic investment can be inferred. In general for real populations energy usage varies markedly depending upon crucial parameters extrinsic to the individual such as resource value and intrinsic ones such as age, and is thus an important factor to consider when modelling.

Potential costs of heterospecific sexual interactions in golden orbweb spiders (Nephila spp.).

Though not uncommon in other animals, heterospecific mating is rarely reported in arachnids. We investigated sexual interactions among four closely related and syntopical African golden orbweb spiders, Nephila inaurata, N. fenestrata, N. komaci, and N. senegalensis. In two South African localities, female webs were often inhabited by heterospecific males that sometimes outnumbered conspecifics. Species association of males with females was random in nature. In subsequent laboratory choice experiments, N. inaurata males chose heterospecific females in 30% of trials. We also observed natural mating interactions between N. inaurata males and N. komaci females, and between N. komaci males and N. inaurata females in laboratory experiments. While heterospecific mating in the laboratory never produced offspring, conspecific mating did. We discuss potential ecological and evolutionary consequences of heterospecific mating interactions in Nephila that may be particularly costly to the rarer species.

The role of ultraviolet colour in the assessment of mimetic accuracy between Batesian mimics and their models: a case study using ant-mimicking spiders.

The use of ultraviolet (UV) cues for intra- and inter-specific communication is common in many animal species. Still, the role of UV signals under some predator-prey contexts, such as Batesian mimicry, is not clear. Batesian mimicry is a defensive strategy by which a palatable species (the mimic) resembles an unpalatable or noxious species (the model) to avoid predation. This strategy has evolved independently in many different taxa that are predated by species capable of UV perception. Moreover, there is considerable variation in how accurately Batesian mimics resemble their models across species. Our aim was to investigate how UV colour contributed to mimetic accuracy using several ant-mimicking spider species as a case study. We measured the reflectance spectrum (300-700 nm) for several species of mimics and models, and we tested whether they differ in visible and UV colour. We modelled whether two different predators could discriminate between mimics and models using colour information. We found that generally, ant-mimicking spiders differed significantly from their ant models in UV colour and that information from the visible range of light cannot be extrapolated into the UV. Our modelling suggested that wasps should be able to discriminate between mimics and models combining information from visible and the UV light, whereas birds may not discriminate between them. Thus, we show that UV colour can influence mimic accuracy and we discuss its potential role in Batesian mimicry. We conclude that colour, especially in the UV range, should be taken into account when measuring mimetic accuracy.

Physiological costs during the first maternal care in the wolf spider Pardosa saltans (Araneae, Lycosidae).

Many arachnids like other terrestrial arthropods, provide extensive maternal care. Few studies have quantified the underlying physiological costs of maternal care. We investigated how maternal care affects the free-moving wolf spider's (Pardosa saltans) energy requirements. We described in detail their basic reproduction biology (i.e. carrying cocoon and young) and we evaluated the variation in the females' energy reserves during maternal care. Our results show that mothers guard eggs until hatching and then guard their spiderlings for 27-30 more days. Laboratory observations indicated that spiderlings start leaving the maternal abdomen gradually 5-7days after hatching. Females carry an egg sac (cocoon) that can weigh up to 77% of their post-reproduction weight and carry young that weigh 87-100% of their body mass. Females lost weight over time despite regular food intake, while carrying cocoon and young; but their weights increased gradually during the dispersal of young. The contributions of proteins, glucose and triglycerides to maintain females' energy were calculated. Their energetic state varied during maternal care, in particular lipid levels declined, during the care of spiderlings when the females' predatory behaviour was inhibited. Our results show that the maternal care provided by P. saltans females is particularly costly physiologically, during the 30days following egg sac formation and development of spiderlings, even when food is available.

Low cost venom extractor based on Arduino(®) board for electrical venom extraction from arthropods and other small animals.

Extracting venom from small species is usually challenging. We describe here an affordable and versatile electrical venom extractor based on the Arduino(®) Mega 2560 Board, which is designed to extract venom from arthropods and other small animals. The device includes fine tuning of stimulation time and voltage. It was used to collect venom without apparent deleterious effects, and characterized for the first time the venom of Zoropsis spinimana, a common spider in French Mediterranean regions.

Selection for costly sexual traits results in a vacant mating niche and male dimorphism.

The expected strong directional selection for traits that increase a male's mating ability conflicts with the frequent observation that within species, males may show extreme variation in sexual traits. These male reproductive polymorphisms are usually attributed to direct male-male competition. It is currently unclear, however, how directional selection for sexually selected traits may convert into disruptive selection, and if female preference for elaborate traits may be an alternative mechanism driving the evolution of male polymorphism. Here, we explore this mechanism using the polyandric dwarf spider Oedothorax gibbosus as a model. We first show that males characterized by conspicuous cephalic structures serving as a nuptial feeding device ("gibbosus males") significantly outperform other males in siring offspring of previously fertilized females. However, significant costs in terms of development time of gibbosus males open a mating niche for an alternative male type lacking expensive secondary sexual traits. These "tuberosus males" obtain virtually all fertilizations early in the breeding season. Individual-based simulations demonstrate a hitherto unknown general principle, by which males selected for high investment to attract females suffer constrained mating opportunities. This creates a vacant mating niche of unmated females for noninvesting males and, consequently, disruptive selection on male secondary sexual traits.

Optimal numbers of matings: the conditional balance between benefits and costs of mating for females of a nuptial gift-giving spider.

In species where females gain a nutritious nuptial gift during mating, the balance between benefits and costs of mating may depend on access to food. This means that there is not one optimal number of matings for the female but a range of optimal mating numbers. With increasing food availability, the optimal number of matings for a female should vary from the number necessary only for fertilization of her eggs to the number needed also for producing these eggs. In three experimental series, the average number of matings for females of the nuptial gift-giving spider Pisaura mirabilis before egg sac construction varied from 2 to 16 with food-limited females generally accepting more matings than well-fed females. Minimal level of optimal mating number for females at satiation feeding conditions was predicted to be 2-3; in an experimental test, the median number was 2 (range 0-4). Multiple mating gave benefits in terms of increased fecundity and increased egg hatching success up to the third mating, and it had costs in terms of reduced fecundity, reduced egg hatching success after the third mating, and lower offspring size. The level of polyandry seems to vary with the female optimum, regulated by a satiation-dependent resistance to mating, potentially leaving satiated females in lifelong virginity.

Animal personality aligns task specialization and task proficiency in a spider society.

Classic theory on division of labor implicitly assumes that task specialists are more proficient at their jobs than generalists and specialists in other tasks; however, recent data suggest that this might not hold for societies that lack discrete worker polymorphisms, which constitute the vast majority of animal societies. The facultatively social spider Anelosimus studiosus lacks castes, but females exhibit either a "docile" or "aggressive" phenotype. Here we observed the propensity of individual females of either phenotype to perform various tasks (i.e., prey capture, web building, parental care, and colony defense) in mixed-phenotype colonies. We then measured the performance outcomes of singleton individuals of either phenotype at each task to determine their proficiencies. Aggressive females participated more in prey capture, web building, and colony defense, whereas docile females engaged more in parental care. In staged trials, aggressive individuals were more effective at capturing prey, constructing webs, and defending the colony, whereas docile females were more effective at rearing large quantities of brood. Thus, individuals' propensity to perform tasks and their task proficiencies appear to be adaptively aligned in this system. Moreover, because the docile/aggressive phenotypes are heritable, these data suggest that within-colony variation is maintained because of advantages gleaned by division of labor.

Individual personalities shape task differentiation in a social spider.

Deciphering the mechanisms involved in shaping social structure is key to a deeper understanding of the evolutionary processes leading to sociality. Individual specialization within groups can increase colony efficiency and consequently productivity. Here, we test the hypothesis that within-group variation in individual personalities (i.e. boldness and aggression) can shape task differentiation. The social spider Stegodyphus sarasinorum (Eresidae) showed task differentiation (significant unequal participation) in simulated prey capture events across 10-day behavioural assays in the field, independent of developmental stage (level of maturation), eliminating age polyethism. Participation in prey capture was positively associated with level of boldness but not with aggression. Body size positively correlated with being the first spider to emerge from the colony as a response to prey capture but not with being the first to attack, and dispersal distance from experimental colonies correlated with attacking but not with emerging. This suggests that different behavioural responses to prey capture result from a complex set of individual characteristics. Boldness and aggression correlated positively, but neither was associated with body size, developmental stage or dispersal distance. Hence, we show that personalities shape task differentiation in a social spider independent of age and maturation. Our results suggest that personality measures obtained in solitary, standardized laboratory settings can be reliable predictors of behaviour in a social context in the field. Given the wealth of organisms that show consistent individual behavioural differences, animal personality could play a role in social organization in a diversity of animals.

Effects of within-colony competition on body size asymmetries and reproductive skew in a social spider.

Reproductive partitioning is a key component of social organization in groups of cooperative organisms. In colonies of permanently social spiders of the genus Stegodyphus less than half of the females reproduce, while all females, including nonreproducers, perform suicidal allo-maternal care. Some theoretical models suggest that reproductive skew is a result of contest competition within colonies, leading to size hierarchies where only the largest females become reproducers. We investigated the effect of competition on within-group body size variation over six months in S. dumicola, by manipulating food level and colony size. We found no evidence that competition leads to increased size asymmetry within colonies, suggesting that contest competition may not be the proximate explanation for reproductive skew. Within-colony body size variation was high already in the juvenile stage, and did not increase over the course of the experiment, suggesting that body size variation is shaped at an early stage. This might facilitate task specialization within colonies and ensure colony-level reproductive output by early allocation of reproductive roles. We suggest that reproductive skew in social spiders may be an adaptation to sociality selected through inclusive fitness benefits of allo-maternal care as well as colony-level benefits maximizing colony survival and production.

Task specialization in two social spiders, Stegodyphus sarasinorum (Eresidae) and Anelosimus eximius (Theridiidae).

Understanding the social organization of group-living organisms is crucial for the comprehension of the underlying selective mechanisms involved in the evolution of cooperation. Division of labour and caste formation is restricted to eusocial organisms, but behavioural asymmetries and reproductive skew is common in other group-living animals. Permanently, social spiders form highly related groups with reproductive skew and communal brood care. We investigated task differentiation in nonreproductive tasks in two permanently and independently derived social spider species asking the following questions: Do individual spiders vary consistently in their propensity to engage in prey attack? Are individual spiders' propensities to engage in web maintenance behaviour influenced by their previous engagement in prey attack? Interestingly, we found that both species showed some degree of task specialization, but in distinctly different ways: Stegodyphus sarasinorum showed behavioural asymmetries at the individual level, that is, individual spiders that had attacked prey once were more likely to attack prey again, independent of their body size or hunger level. In contrast, Anelosimus eximius showed no individual specialization, but showed differentiation according to instar, where adult and subadult females were more likely to engage in prey attack than were juveniles. We found no evidence for division of labour between prey attack and web maintenance. Different solutions to achieve task differentiation in prey attack for the two species studied here suggest an adaptive value of task specialization in foraging for social spiders.

Spinning a laser web: predicting spider distributions using LiDAR.

LiDAR remote sensing has been used to examine relationships between vertebrate diversity and environmental characteristics, but its application to invertebrates has been limited. Our objectives were to determine whether LiDAR-derived variables could be used to accurately describe single-species distributions and community characteristics of spiders in remote forested and mountainous terrain. We collected over 5300 spiders across multiple transects in the Bavarian National Park (Germany) using pitfall traps. We examined spider community characteristics (species richness, the Shannon index, the Simpson index, community composition, mean body size, and abundance) and single-species distribution and abundance with LiDAR variables and ground-based measurements. We used the R2 and partial R2 provided by variance partitioning to evaluate the predictive power of LiDAR-derived variables compared to ground measurements for each of the community characteristics. The total adjusted R2 for species richness, the Shannon index, community species composition, and body size had a range of 25-57%. LiDAR variables and ground measurements both contributed >80% to the total predictive power. For species composition, the explained variance was approximately 32%, which was significantly greater than expected by chance. The predictive power of LiDAR-derived variables was comparable or superior to that of the ground-based variables for examinations of single-species distributions, and it explained up to 55% of the variance. The predictability of species distributions was higher for species that had strong associations with shade in open-forest habitats, and this niche position has been well documented across the European continent for spider species. The similar statistical performance between LiDAR and ground-based measures at our field sites indicated that deriving spider community and species distribution information using LiDAR data can provide not only high predictive power at relatively low cost, but may also allow unprecedented mapping of community- and species-level spider information at scales ranging from stands to landscapes. Therefore, LiDAR is a viable tool to assist species-specific conservation as well as broader biodiversity planning efforts not only for a growing list of vertebrates, but for invertebrates as well.

The adaptive value of gluttony: predators mediate the life history trade-offs of satiation threshold.

Animals vary greatly in their tendency to consume large meals. Yet, whether or how meal size influences fitness in wild populations is infrequently considered. Using a predator exclusion, mark-recapture experiment, we estimated selection on the amount of food accepted during an ad libitum feeding bout (hereafter termed 'satiation threshold') in the wolf spider Schizocosa ocreata. Individually marked, size-matched females of known satiation threshold were assigned to predator exclusion and predator inclusion treatments and tracked for a 40-day period. We also estimated the narrow-sense heritability of satiation threshold using dam-on-female-offspring regression. In the absence of predation, high satiation threshold was positively associated with larger and faster egg case production. However, these selective advantages were lost when predators were present. We estimated the heritability of satiation threshold to be 0.56. Taken together, our results suggest that satiation threshold can respond to selection and begets a life history trade-off in this system: high satiation threshold individuals tend to produce larger egg cases but also suffer increased susceptibility to predation.

Discovery of the largest orbweaving spider species: the evolution of gigantism in Nephila.

BACKGROUND: More than 41,000 spider species are known with about 400-500 added each year, but for some well-known groups, such as the giant golden orbweavers, Nephila, the last valid described species dates from the 19(th) century. Nephila are renowned for being the largest web-spinning spiders, making the largest orb webs, and are model organisms for the study of extreme sexual size dimorphism (SSD) and sexual biology. Here, we report on the discovery of a new, giant Nephila species from Africa and Madagascar, and review size evolution and SSD in Nephilidae. METHODOLOGY: We formally describe N. komaci sp. nov., the largest web spinning species known, and place the species in phylogenetic context to reconstruct the evolution of mean size (via squared change parsimony). We then test female and male mean size correlation using phylogenetically independent contrasts, and simulate nephilid body size evolution using Monte Carlo statistics. CONCLUSIONS: Nephila females increased in size almost monotonically to establish a mostly African clade of true giants. In contrast, Nephila male size is effectively decoupled and hovers around values roughly one fifth of female size. Although N. komaci females are the largest Nephila yet discovered, the males are also large and thus their SSD is not exceptional.

Costs and benefits of freezing behaviour in the harvestman Eumesosoma roeweri (Arachnida, Opiliones).

Animals present an enormous variety of behavioural defensive mechanisms, which increase their survival, but often at a cost. Several animal taxa reduce their chances of being detected and/or recognized as prey items by freezing (remaining completely motionless) in the presence of a predator. We studied costs and benefits of freezing in immature Eumesosoma roeweri (Opiliones, Sclerosomatidae). Preliminary observations showed that these individuals often freeze in the presence of the syntopic predatory spider Schizocosa ocreata (Araneae, Lycosidae). We verified that harvestmen paired with predators spent more time freezing than when alone or when paired with a conspecific. Then, we determined that predator chemical cues alone did not elicit freezing behaviour. Next, we examined predator behaviour towards moving/non-moving prey and found that spiders attacked moving prey significantly more, suggesting an advantage of freezing in the presence of a predator. Finally, as measure of the foraging costs of freezing, we found that individuals paired with a predator for 2h gained significantly less weight than individuals paired with a conspecific or left alone. Taken together, our results suggest that freezing may protect E. roeweri harvestmen from predatory attacks by wolf spiders, but at the cost of reduced food and/or water intake.

Undersampling bias: the null hypothesis for singleton species in tropical arthropod surveys.

1. Frequency of singletons - species represented by single individuals - is anomalously high in most large tropical arthropod surveys (average, 32%). 2. We sampled 5965 adult spiders of 352 species (29% singletons) from 1 ha of lowland tropical moist forest in Guyana. 3. Four common hypotheses (small body size, male-biased sex ratio, cryptic habits, clumped distributions) failed to explain singleton frequency. Singletons are larger than other species, not gender-biased, share no particular lifestyle, and are not clumped at 0.25-1 ha scales. 4. Monte Carlo simulation of the best-fit lognormal community shows that the observed data fit a random sample from a community of approximately 700 species and 1-2 million individuals, implying approximately 4% true singleton frequency. 5. Undersampling causes systematic negative bias of species richness, and should be the default null hypothesis for singleton frequencies. 6. Drastically greater sampling intensity in tropical arthropod inventory studies is required to yield realistic species richness estimates. 7. The lognormal distribution deserves greater consideration as a richness estimator when undersampling bias is severe.

Survival benefits select for group living in a social spider despite reproductive costs.

The evolution of cooperation requires benefits of group living to exceed costs. Hence, some components of fitness are expected to increase with increasing group size, whereas others may decrease because of competition among group members. The social spiders provide an excellent system to investigate the costs and benefits of group living: they occur in groups of various sizes and individuals are relatively short-lived, therefore life history traits and Lifetime Reproductive Success (LRS) can be estimated as a function of group size. Sociality in spiders has originated repeatedly in phylogenetically distant families and appears to be accompanied by a transition to a system of continuous intra-colony mating and extreme inbreeding. The benefits of group living in such systems should therefore be substantial. We investigated the effect of group size on fitness components of reproduction and survival in the social spider Stegodyphus dumicola in two populations in Namibia. In both populations, the major benefit of group living was improved survival of colonies and late-instar juveniles with increasing colony size. By contrast, female fecundity, female body size and early juvenile survival decreased with increasing group size. Mean individual fitness, estimated as LRS and calculated from five components of reproduction and survival, was maximized for intermediate- to large-sized colonies. Group living in these spiders thus entails a net reproductive cost, presumably because of an increase in intra-colony competition with group size. This cost is traded off against survival benefits at the colony level, which appear to be the major factor favouring group living. In the field, many colonies occur at smaller size than expected from the fitness curve, suggesting ecological or life history constraints on colony persistence which results in a transient population of relatively small colonies.

A total evidence assessment of the phylogeny of North American euctenizine trapdoor spiders (Araneae, Mygalomorphae, Cyrtaucheniidae) using Bayesian inference.

North American trapdoor spiders of the subfamily Euctenizinae (Cyrtaucheniidae) are among the most diverse mygalomorph spiders (trapdoor spiders, tarantulas, and their relatives) on the continent in terms of species numbers and ecological habits. We present a generic level phylogenetic study of the subfamily based on a total evidence approach. Our dataset comprises approximately 3.7 kb of molecular characters (18S and 28S rRNA gene sequences) and 71 morphological characters scored for 32 taxa. When analyzed independently, these data sets, particularly the morphology, depict very different views of mygalomorph and euctenizine relationships, albeit with weak support. However, when these data are combined we recover a tree topology that is supported by high posterior probability for most nodes. The combined data recover a phylogenetic pattern for euctenizines different than previously published and indicate the presence of a narrowly endemic new genus from central California. While euctenizine monophyly is unequivocal, the monophyly of a number of other mygalomorph groups is questionable (e.g., Cyrtaucheniidae, Mecicobothriodina, Rastelloidina). This non-monophyly is noteworthy, as our analysis represents the first employing a total evidence approach for mygalomorphs, a group known to be morphologically conservative.